Chondrichthyes: Neoselachii) in the Jurassic of Normandy
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FIRST MENTION OF THE FAMILY PSEUDONOTIDANIDAE (CHONDRICHTHYES: NEOSELACHII) IN THE JURASSIC OF NORMANDY by Gilles CUNY (1) and Jérôme TABOUELLE (2) ABSTRACT The discovery of a tooth of cf. Pseudonotidanus sp. is reported from the Bathonian of Normandy. Its morphology supports the transfer of the species terencei from the genus Welcommia to the genus Pseudonotidanus. It also supports the idea that Pseudonotidanidae might be basal Hexanchiformes rather than Synechodontiformes. KEYWORDS Normandy, Arromanches, Jurassic, Bathonian, Chondrichthyes, Elasmobranchii, Synechodontiformes, Hexanchiformes, Pseudonotidanidae. RÉSUMÉ La découverte d’une dent de cf. Pseudonotidanus sp. est signalée dans le Bathonien de Normandie. Sa morphologie soutient le transfert de l’espèce terencei du genre Welcommia au genre Pseudonotidanus . Elle soutient également l’idée que les Pseudonotidanidae pourraient appartenir aux Hexanchiformes basaux plutôt qu’aux Synechodontiformes. MOTS-CLEFS Normandie, Arromanches, Jurassique, Bathonien, Chondrichthyes, Elasmobranchii, Synechodontiformes, Hexanchiformes, Pseudonotidanidae. References of this article: CUNY G. and TABOUELLE J. (2014) – First mention of the family Pseudonotidanidae (Chondrichthyes: Neoselachii) in the Jurassic of Normandy. Bulletin Sciences et Géologie Normandes , tome 7, p. 21-28. 1 – INTRODUCTION In 2004, Underwood and Ward erected the new family Pseudonotidanidae for sharks showing teeth with a hexanchid-like crown (compressed labio-lingually with several cusps) and a Palaeospinacid-like root (lingually expanded and flat based), hence offering an opportunity to better understand the phylogenetic relationships between the Hexanchiformes and the Synechodontiformes, which might be closely related (Duffin & Ward, 1993; Underwood, 2006). ____________________ (1) Université Claude Bernard Lyon 1, Campus de la Doua, Bâtiment Géode 2, rue Raphaël Dubois 69622 VILLEURBANNE CEDEX, France Email : [email protected] (2) Museum Elbeuf – La Fabrique des Savoirs 7, cours Gambetta 76500 ELBEUF, France Email : [email protected] - 21 - Gilles CUNY and Jérôme TABOUELLE Pseudonotidanidae were first included into Synechodontiformes (Underwood & Ward, 2004), an idea accepted by Klug & Kriwet (2008, 2010), Klug et al . (2009), Klug (2010) and Kriwet & Klug (2011), but this family may as well belongs to the Hexanchiformes according to other authors (Underwood, 2006; Cappetta, 2012; Maisey, 2012). Based on enameloid microstructure, Guinot & Cappetta (2011) also considered Welcommia to be a Hexanchiformes. So far, the family Pseudonotidanidae is represented by two genera, encompassing three and two species respectively ( Pseudonotidanus politus, P. semirugosus, P. terencei, Welcommia bodeuri, W. cappettai). Cappetta (2012) considers indeed that the species terencei is better attributed to the genus Pseudonotidanus than to Welcommia. The genus Pseudonotidanus has been reported from the Toarcian of Germany and Belgium, the Bathonian and Callovian of England and the Oxfordian of France, whereas the genus Welcommia is known from the Oxfordian of Germany and Valanginian of Southern France (Underwood & Ward, 2004; Klug and Kriwet, 2010; Cappetta, 2012). We add here a report of this family in the Bathonian of Normandy. 2 – GEOLOGICAL SETTING The specimen, housed in the collection of Elbeuf Museum with the number 2013.5.1 ME, was surface collected on the intertidal zone at Arromanches (Fig. 1), at the top of the lower member of the Port-en-Bessin Marls Formation ( Tenuiplicatus zone, Lower Bathonian, Arromanches 2 surface). To the East of Port-en-Bessin, the rocky intertidal zone is made of the last beds of the Porifera Limestone (Calcaire à spongiaires). In the background, the cliff is made of three parts. The basal part is a vertical cliff made of a series of greyish marlstone, the Port-en-Bessin Marls (Marnes de Port-en-Bessin). The middle part is a gently sloping scree measuring 25 m and showing sediment slides with some marls outcropping. This slope is covered by vegetation. Near the top, the cliff becomes stiff again and is made of a series of marls and limestones, the limestone beds becoming thicker to make the massive yellowish limestone, called Saint-Pierre-du-Mont Limestone (Calcaire de Saint-Pierre-du-Mont). The stratigraphic log of the cliff (log of the « Tour Vauban ») is thus as follows: 1. Saint-Pierre-du-Mont Limestone (12 m): Middle Bathonian. 2. Port-en-Bessin Marls (35-40 m): Lower to Middle Bathonian. 3. Porifera Limestones (outcropping thickness: 2 m): Upper Bajocian. Fig. 1: Geographical location of Arromanches (red star) (France, Calvados) (Dugué et al ., 1998). - 22 - FIRST MENTION OF THE FAMILY PSEUDONOTIDANIDAE Only the upper part of the Porifera Limestone and the lower part of Port-en-Bessin Marls are easily accessible. A detailed log of the Bajocian/Bathonian transition has been published by Rioult & Fily (1975, 1980) and Rioult et al . (1991) (Fig. 2). Fig. 2: Log and sequence stratigraphy of the eastern cliff at Port-en-Bessin (Lower and Middle Bathonian). The red arrow marks the Arromanches 2 surface, where 2013.5.1 ME was found. From Rioult et al. (1991). The top of the Porifera Limestone is made of a white limestone with lumps of clay and pyrite, the latter often hydrated into limonite. It is bioturbated and ends with an erosion surface, the Port-en- Bessin 1 surface. This surface is overlain by three thin beds of hard, argillaceous limestone, called beds a, b and c of the transition zone (Guillaume, 1927), which makes the transition between the Bajocian Porifera limestone and the Bathonian Port-en-Bessin Marls. Each of these beds corresponds to one of the three sub-zones of the Zigzag zone (Lower Bathonian). The Port-en-Bessin Marls ( Tenuiplicatus and Progracilis zones, Lower Bathonian to lowermost Middle Bathonian) are made of a series of grey marls and argillaceous limestones that outcrop in many places around the Bessin area, but more particularly in the coastal cliffs between Grandcamp and Arromanches (Fig. 3). Three units are distinguished in the Port-en-Bessin Marls (Rioult & Fily, 1975 ; Fily et al. , 1989), but only the lower one (10 to 12 m, Tenuiplicatus zone, Lower Bathonian) is accessible in the cliffs. It is made of a series of decimetric marly beds grading into argillaceous limestone. The latter appears often discontinuous, without a clear upper boundary. Their thickness and coarseness raise regularly through two parasequences, reflecting the progradation of a shelf prism. The upper boundary of this unit is marked by two beds of limestone (0.5 m thick at Port-en-Bessin, 1 m thick at Arromanches) with more bioturbations, often overhanging in the cliffs. These two beds are interpreted as a thin transgressive interval. This lithology marker holds at its base the transgressive surface Arromanches 1 and at its top the Arromanches 2 surface, a sedimentary discontinuity characterized by mineralogical changes, where smectite replaces kaolinite, as well as by faunistic changes, with the disappearance of the endofauna and explosion of plankton. It marks the maximum flooding surface (MFS) and the boundary between the Lower and Middle Bathonian. - 23 - Gilles CUNY and Jérôme TABOUELLE Fig. 3: The cliff at Arromanches, displaying the Lower and Middle Bathonian Port-en-Bessin Marls with the Arromanches 1 and Arromanches 2 surfaces. 3 – DESCRIPTION Class Chondrichthyes Huxley, 1880 Subclass Elasmobranchii Bonaparte, 1838 Subcohort Neoselachii Compagno, 1977 Order ?Hexanchiformes Buen, 1926 Family Pseudonotidanidae Underwood & Ward, 2004 Genus Pseudonotidanus Underwood & Ward, 2004 cf. Pseudonotidanus sp. Material: 1 tooth (2013.5.1. ME, fig. 4) The tooth 2013.5.1 ME measures 10 mm mesio-distally, 4 mm labio-lingually and is 8 mm high. It is almost complete, although the mesial extremity of the tooth is missing. It shows a main cusp flanked by 6 cusplets distally, the sixth one being incipient and 5 cusplets mesially. The main cusp and first distal cusplets are inclined distally. The distal cusplets are larger than the mesial ones. They show, like the main cusp convex labial and lingual faces whereas the mesial ones are more flattened. Lateral cusplets decrease regularly in size towards the mesial and distal extremities of the crown. The first and third mesial cusplets show a small extra cusplet mesially. There is a well developed cutting edge, which is continuous from the main cusp to the most distal and mesial cusplets. The enameloid is smooth on both sides of the crown, but there is a band devoid of enameloid at the base of the lingual face of the crown (lingual neck, fig. 4C). - 24 - FIRST MENTION OF THE FAMILY PSEUDONOTIDANIDAE Fig. 4: Tooth of cf. Pseudonotidanus sp. from Arromanches in A: labial, B: distal, C: lingual, and D: basal views. The root is quite deep in labial view and shows foramina mainly restricted to its lower part (Fig. 4A). There are three enlarged foramina at the base, just below the main cusp, with a partially open basal wall, thus forming three short grooves on the labial part of the basal face of the root (Fig. 4D). The root is slightly projected lingually with a concave face showing a row of slit-like foramina in its middle part. There is also one enlarged foramen in the basal part, just below the main cusp. It is flanked distally by a rather large, slit-like foramen, again lacking part of its basal wall. The basal face of the root is lozenge-shaped and show many foramina randomly distributed. Its basal face forms with the axis of the crown